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Copepods are marine invertebrates with a key role at the basis of marine food webs due to their high biomass as well as their elevated fatty acid (FA) content, particularly eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA), two FA which have a well demonstrated role in growth and reproduction in marine organisms. Temperature is a key driver of organisms fitness as well as ecosystem functioning and sea surface temperature is expected to rise under all CO2 emission scenarios. Thus, understanding how copepods will response to such changes is crucial given their role in marine food webs. While the majority of research has focused on planktonic copepod species, less is known for benthic species, particularly non-model species. In this study we expose Microarthridion littorale, an intertidal benthic copepod to a temperature gradient (12, 15, 18, 21 and 24 °C) including current environmental variability as well as future scenarios. Individuals were retrieved from a mudflat in the Westerschelde estuary (51°21′ 24′′ N, 3° 42′ 51′′E) and cultured at lab conditions. Survival and FA were measured after 18 days exposure. Growth rates and nauplii production were also measured for each temperature treatment. We found decreased survival, EPA and DHA content with increased temperature as well as increased growth rates and detrimental effects for nauplii production. Benthic copepods are prey for many juvenile fish, any changes in their biomass as well as their FA composition in response to temperature changes could therefore amplify to higher trophic levels with important consequences for food web functioning.

Highlights: • Effects of microplastic on marine biota reflect the quality of experimental research. • The quality of published experiments can be quantified from an “ideal” experiment. • Previously published experiments have significantly deviated from “ideal”. • Implementation of proposed criteria can improve future microplastic experiments. Abstract: This article presents a novel conceptual blueprint for an ‘ideal’, i.e., ecologically relevant, microplastic effect study. The blueprint considers how microplastics should be characterized and applied in laboratory experiments, and how biological responses should be measured to assure unbiased data that reliably reflect the effects of microplastics on aquatic biota. This ‘ideal’ experiment, although practically unachievable, serves as a backdrop to improve specific aspects of experimental research on microplastic effects. In addition, a systematic and quantitative literature review identified and quantified departures of published experiments from the proposed ‘ideal’ design. These departures are related mainly to the experimental design of microplastic effect studies failing to mimic natural environments, and experiments with limited potential to be scaled-up to ecosystem level. To produce a valid and generalizable assessment of the effect of microplastics on biota, a quantitative meta-analysis was performed that incorporated the departure of studies from the ‘ideal’ experiment (a measure of experimental quality) and inverse variance (a measure of the study precision) as weighting coefficients. Greater weights were assigned to experiments with higher quality and/or with lower variance in the response variables. This double-weighting captures jointly the technical quality, ecological relevance and precision of estimates provided in each study. The blueprint and associated meta-analysis provide an improved baseline for the design of ecologically relevant and technically sound experiments to understand the effects of microplastics on single species, populations and, ultimately, entire ecosystems.

Ocean warming and acidification affect species populations, but how interactions within communities are affected and how this translates into ecosystem functioning and resilience remain poorly understood. Here we demonstrate that experimental ocean warming and acidification significantly alters the interaction network among porewater nutrients, primary producers, herbivores and burrowing invertebrates in a seafloor sediment community, and is linked to behavioural plasticity in the clam Scrobicularia plana. Warming and acidification induced a shift in the clam’s feeding mode from predominantly suspension feeding under ambient conditions to deposit feeding with cascading effects on nutrient supply to primary producers. Surface-dwelling invertebrates were more tolerant to warming and acidification in the presence of S. plana, most probably due to the stimulatory effect of the clam on their microalgal food resources. This study demonstrates that predictions of population resilience to climate change require consideration of non-lethal effects such as behavioural changes of key species. Changes in ocean temperature and pH will impact on species, as well as impacting on community interactions. Here warming and acidification cause a clam species to change their feeding mode, with cascading effects for the marine sedimentary food web.

Intertidal polychaete aggregations may be protected in the European Union under the Habitats Directive framework as reef habitats. Remote reef mapping remains challenging due to severe and dynamic conditions, as well as cover and spatial resolution requirements. This study (1) evaluated kite aerial photography and low-altitude digital photogrammetry to map and monitor intertidal aggregations of a sessile tube-building polychaete, Lanice conchilega (L. conchilega), and (2) developed a protocol for its remote identification. Monthly campaigns yielded 12 aerial image sets which were processed using structure-from-motion into high-precision digital terrain models and orthophoto mosaics. Maximum likelihood classification distinguished L. conchilega from bare sediment with an accuracy of 70% ± 23.2%. Aggregations were delineated by extracting elements of positive elevation from local difference models. The method has proven useful to detect high-value aggregations, distinguishing these consistently. Nevertheless, systematic biases were present during delineation, and further characterisation of reference aggregations may improve detection.

Soft-sediment biogeochemistry is influenced by the bioturbation activity of benthic invertebrates. We investigated whether the effect of two macrobenthos bioturbators, Limecola balthica and Hediste diversicolor, on sediment oxygen uptake can be described by allometric principles of metabolic activity scaling with animal body size and population biomass. Microcosms containing reconstructed populations to control density and individual body size were used to compare bioturbation effects and allometric scaling principles between a sandy and muddy sediment. Both species facilitated oxygen uptake in both sediment types, and a major portion of the variance in sediment metabolism (60–98%) could be explained by the per capita body size and density, and total population biomass. The allometric relationship with the stimulated sediment metabolism was similar in sand and mud for Hediste and strongly related to the increasing burrow ventilation rate with population biomass. Limecola irrigated less in mud but stimulated sediment metabolism more in mud in comparison to in sand. We discuss how physico-chemical differences between both sediment types, possible changes in activity, and size-dependent irrigation dynamics can explain the variable effects of Limecola on sediment metabolism. Overall, we provide empirical evidence that allometric laws can be used to upscale bioturbation effects on ecosystem functioning in marine soft sediments from the individual to the population level.

The presence and behaviour of bivalves can affect the functioning of seafloor sediments through the irrigation of deeper strata by feeding and respiring through siphonal channels. Here, we investigated the physiological response and consecutive impact on functioning and body condition of the white furrow shell Abra alba in three pH treatments (pH = 8.2, pH = 7.9 and pH = 7.7). Although no pH effect on survival was found, lowered respiration and calcification rates, decreased energy intake (lower absorption rate) and increased metabolic losses (increased excretion rates) occurred at pH ∼ 7.7. These physiological responses resulted in a negative Scope for Growth and a decreased condition index at this pH. This suggests that the physiological changes may not be sufficient to sustain survival in the long term, which would undoubtedly translate into consequences for ecosystem functioning.

The distinct spatial variability in microplastic concentrations between marine regions and habitats calls for a better understanding about the transport pathways of this omnipresent pollutant in the marine environment. This study provides empirical evidence that a sessile filter feeder, the Blue mussel M. edulis, accelerates microplastic deposition by aggregating them into sinking particulate faeces and pseudofaeces. After settling to the seafloor, the bioturbation of benthic fauna quickly buries these microplastics. Collectively, these results suggest that if such biologically-mediated benthic-pelagic coupling would be integrated into hydrodynamic transport models, the spatial variability and source-sink dynamics of microplastics would be better understood. It is proposed that microplastic pollution is monitored through sampling that takes into account faeces and pseudofaeces underneath filter feeders. The implications of this detrital pathway for microplastic transfer to the seafloor, and the role of shellfish mariculture in this process, are discussed. Studies that consider filter feeders and benthic communities from other regions, and during different seasons, are needed to validate the proposed biological pump mechanism across space and time.

Our study presents a revision of the structural characteristics of the soft-bottom benthic communities in the Belgian Part of the North Sea (BPNS) and provides, for the first time, an insight into their functional characteristics. Compared to previous research, the dataset expanded considerably on a spatial and temporal scale (1994–2012). Five communities were identified, each with their own structural characteristics, indicator species, sediment properties and spatial distribution. The offshore area was dominated by the Hesionura elongata community (formerly Ophelia borealis), and although previously described by low diversity and abundance, this study shows that this community has the second highest sample diversity and moderate densities. Also, a new community, the Magelona-Ensis leei community, was found in very shallow water and characterized by low diversity and the dominance of the non-indigenous species E. leei. The structural characteristics and spatial distribution of the previously described Limecola balthica, Abra alba and Nephtys cirrosa community remained largely unchanged. Though structurally distinct, the communities overlap in some of their functional attributes. Analysis of biological traits revealed that in the coarser permeable sands, both the N. cirrosa and H. elongata community harbor more free living mobile individuals, causing diffusive mixing. The finer sand (A. alba, Magelona- Ensis leei) and muddy (L. balthica) communities hold more sessile, tube building and burrow dwelling species. Apart from diffusive mixing, sediment reworking in these finer sands occurs by surface deposition and downward convection. In combination with higher densities and biomass, this results in a higher bioturbation potential (BPc) for the Abra alba and Magelona-Ensis leei community. Finally, measures for functional diversity and redundancy were calculated, and were also highest in the Abra alba and Hesionura elongata community, thus demonstrating the importance of both communities for conservation purposes from a structural as well as a functional perspective. The analyses revealed that even in well-studied areas new data give new insights, with important consequences for management. Our study forms a benchmark for future ecosystem health status assessments in the framework of international and national nature management obligations such as the European Marine Strategy Framework Directive (MSFD), the Habitats Directive and the Marine Spatial Planning Directive.

In future global change scenarios the surface ocean will experience continuous acidification and rising temperatures. While effects of both stressors on marine, benthic communities are fairly well studied, consequences of the interaction of both factors remain largely unknown. We performed a short-term microcosm experiment exposing a soft-bottom community from an intertidal flat in the Westerscheldt estuary to two levels of seawater pH (ambient pHT = 7.9, reduced pHT = 7.5) and temperature (10 °C ambient and 13 °C elevated temperature) in a crossed design. After 8 weeks, meiobenthic community structure and nematode staining ratios, as a proxy for mortality, were compared between treatments and structural changes were related to the prevailing abiotic conditions in the respective treatments (pore water pHT, sediment grain size, total organic matter content, total organic carbon and nitrogen content, phytopigment concentrations and carbonate concentration). Pore water pHT profiles were significantly altered by pH and temperature manipulations and the combination of elevated temperature and reduced pH intensified the already more acidic porewater below the oxic zone. Meiofauna community composition was significantly affected by the combination of reduced pH and elevated temperature resulting in increased densities of predatory Platyhelminthes, reduced densities of Copepoda and Nauplii and complete absence of Gastrotricha compared to the experimental control. Furthermore, nematode staining ratio was elevated when seawater pH was reduced pointing towards reduced degradation rates of dead nematode bodies. The observed synergistic interactions of pH and temperature on meiobenthic communities and abiotic sediment characteristics underline the importance of multistressor experiments when addressing impacts of global change on the marine environment.

Salt marshes in urban watersheds are prone to microplastics (MP) pollution due to their hydrological characteristics and exposure to urban runoff, but little is known about MP distributions in species from these habitats. In the current study, MP occurrence was determined in six benthic invertebrate species from salt marshes along the North Adriatic lagoons (Italy) and the Schelde estuary (Netherlands). The species represented different feeding modes and sediment localisation. The 96% of the analysed specimens (330) did not contain any MP, which was consistent across different regions and sites. Suspension and facultative deposit-feeding bivalves exhibited a lower MP occurrence (0.5-3%) relative to omnivores (95%) but contained a much more variable distribution of MP sizes, shapes and polymers. The study provides indications that MP physicochemical properties and species' ecological traits could all influence MP exposure, uptake and retention in benthic organisms inhabiting European salt marsh ecosystems.

Functional trait approaches advance the understanding of biodiversity–ecosystem function (BDEF) relationships and its control by the environmental context. Application of these insights into management remains constrained due to lack of evidence from real-world ecosystems that capture the natural spatial and temporal gradients at which biodiversity and environmental conditions operate. In this study we measured macrofauna community traits, ecosystem processes and abiotic properties at 9 locations during 4 months, spanning a wide gradient in sedimentary habitats and salinity in the Scheldt estuary, and quantified the (a)biotic contribution to sediment community oxygen consumption, as a measure of ecosystem function. We found that functional attributes of the macrofauna community and its effect on bio-irrigation can predict ecosystem function, but especially during the colder period of the year. This result highlights that generalizations about BDEF relationships, and biodiversity loss on ecosystem functions, are limited whenever this temporal component is not acknowledged.

Litter is omnipresent in the ocean where it can be ingested by marine biota. Although ingestion of microplastics (MPs) is abundantly reported, insights into how MP can influence predator-prey interactions currently limits our understanding of the ecological impact of MPs. Here we demonstrate trophic transfer of MPs from zooplankton to benthic filter feeders, through consumption of contaminated prey (i.e. prey with ingested MP). However, predation rates of contaminated prey were significantly lower as compared to predation rates of prey that had no MPs ingested. As filter feeder clearance rates were not affected by consumption of MPs, the lower predation rates of contaminated prey appear to be primarily explained by disruption in zooplankton swimming behaviour that reduces their filtration risk. This is the first study that shows how MPs can change predator-prey interactions that are involved in the coupling between the pelagic and seabed habitat.

Saltmarshes have been depleted historically, and cumulative stressors threaten their future persistence. We examined experimentally how nutrient availability (high vs. low) affects the responses of Spartina maritima to increased inundation in two mineral soil types (low vs. medium organic). Increased inundation, one of the effects of accelerated sea level rise, had negative effects on most plant growth parameters, but the magnitude varied with soil and nutrient levels, and between plants from different locations. Average differences between inundation treatments were largest at high nutrient conditions in low organic matter soils. We conclude that saltmarsh vegetation would be more drastically affected by increased inundation in low than in medium organic matter soils, and especially in estuaries already under high nutrient availability. This knowledge enhances the prediction of changes at the foreshore of saltmarshes related to sea level rise, and the development of site-specific conservation strategies.

Motivation The BioTIME database contains raw data on species identities and abundances in ecological assemblages through time. These data enable users to calculate temporal trends in biodiversity within and amongst assemblages using a broad range of metrics. BioTIME is being developed as a community-led open-source database of biodiversity time series. Our goal is to accelerate and facilitate quantitative analysis of temporal patterns of biodiversity in the Anthropocene. Main types of variables included The database contains 8,777,413 species abundance records, from assemblages consistently sampled for a minimum of 2 years, which need not necessarily be consecutive. In addition, the database contains metadata relating to sampling methodology and contextual information about each record. Spatial location and grain BioTIME is a global database of 547,161 unique sampling locations spanning the marine, freshwater and terrestrial realms. Grain size varies across datasets from 0.0000000158 km2 (158 cm2) to 100 km2 (1,000,000,000,000 cm2). Time period and grain BioTIME records span from 1874 to 2016. The minimal temporal grain across all datasets in BioTIME is a year. Major taxa and level of measurement BioTIME includes data from 44,440 species across the plant and animal kingdoms, ranging from plants, plankton and terrestrial invertebrates to small and large vertebrates. Software format .csv and .SQL., Global Ecology and Biogeography, 27 (7), ISSN:1466-822X, ISSN:1466-8238

Soft-bottom macrobenthic communities are an essential part of the marine ecosystem for which a healthy status is aimed at. Therefore, insights in the structural and functional (trait based) characteristics of the soft-bottom benthic communities in the Southern part of the North Sea are obtained based on 18 years of data, with a large spatial coverage. The communities show clear differences in their structural properties. The fine sand Abra alba community and coarse sand Hesionura elongata community have the highest species richness and diversity values. The muddy Limecola balthica community and medium sand Nephtys cirrosa community the lowest. However, looking at biological trait characteristics, the communities show some functional redundancy. In the relative coarser, permeable sands, more free living, mobile species, were found causing diffusive mixing, whilst the finer sand and mud communities have more sessile, tube building and burrow dwelling species. With their contribution to bioturbation and bio-irrigation processes, the benthic fauna prove to be essential for the biogeochemical status of these finer sand sediments. Based on this integrated evaluation of biodiversity and functional related characteristics, we will demonstrate how this improve status assessments and an appropriate determination of the habitat sensitivity to different human induced pressures.

The loss or decline in population size of key species can instigate a cascade of effects that have implications for interacting species, therewith impacting biodiversity and ecosystem functioning. We examined how top–down and bottom–up interactions may mediate knock-on effects of a coastal deposit-feeding clam, Macomona liliana (hereafter Macomona), on sandflat meiobenthos densities. Therefore we manipulated densities of Macomona in combination with predator exclusion and experimental shading that was expected to alter microphytobenthos biomass. We show that Macomona regulated densities of meiobenthic (38–500 μm) nematodes, copepods, polychaetes, turbellarians, and ostracodes during the three months of incubation via indirect mechanisms. Predator pressure on Macomona by eagle rays (Myliobatis tenuicaudatus) was found to have a negative effect on densities of some meiobenthic taxa. Furthermore, experimental shading resulted in the loss of a positive relation between Macomona and microphytobenthos biomass, while concurrently increasing the density of some meiobenthic taxa. We suggest that this observation can be explained by the release from bioturbation interference effects of the cockle Austrovenus stutchburyi that was found to thrive in the presence of Macomona under non-shaded conditions. Our results highlight the importance of interactions between macrofaunal bioturbation, microphyte biomass, sediment stability, and predation pressure for the structuring of benthic communities. This experiment illustrates that manipulative field experiments may be particularly suitable to study such multiple indirect mechanisms that regulate ecosystem diversity and related functioning because such approaches may best capture the complex feedbacks and processes that determine ecosystem dynamics.

Anthropogenic CO2 emissions are rapidly changing seawater temperature, pH and carbonate chemistry. This study compares the embryonic development under high pCO2 conditions across the south-north distribution range of the marine clam Limecola balthica in NW Europe. The combined effects of elevated temperature and reduced pH on hatching success and size varied strongly between the three studied populations, with the Gulf of Finland population appearing most endangered under the conditions predicted to occur by 2100. These results demonstrate that the assessment of marine faunal population persistence to future climatic conditions needs to consider the interactive effects of co-occurring physico-chemical alterations in seawater within the local context that determines population fitness, adaptation potential and the system resilience to environmental change.

Mudflats provide a suitable habitat for a functionally diverse invertebrate macrofauna community that can cope with the high variability in sediment physico-chemical conditions associated with the tidal regime. The most common macrofaunal taxonomic groups are annelids, molluscs and crustaceans. Macrofauna are usually the link from primary producers, meiofauna, and detritus to higher trophic levels like wading birds and epibenthic fish that forage in mudflats. Behavioral activities related to macrofauna feeding, burrowing and respiration alter biogeochemistry and mudflat sediment dynamics and are thus of paramount importance for the cycling of energy and matter in mudflats from estuaries, mangroves and coastal lagoons. Variability in abiotic and biotic interactions structures macrofauna communities in space and time and hence defines the influence macrofauna has on the diversity and functioning of the wider ecosystem through direct and indirect interactions.

Ecosystem engineers are organisms that modulate natural resources enabling the survival of other species. They drive environmental change and contribute to several coastal functional attributes such as landscape heterogeneity, sedimentary processes, and coastal protection. Our study focuses on the case of Lanice conchilega, a tube-building ecosystem engineer whose aggregations impact sedimentary processes. This polychaete forms biogenic tube aggregations distributed on the coasts of the northern hemisphere from the shallow intertidal to depths of 1900 m. The aggregations engineer sedimentary processes autogenically by altering water flow at the benthic-boundary layer, and harbor highly diverse infaunal communities as a consequence. This study evaluates the relationships between intertidal L. conchilega aggregations and sedimentary processes at the intertidal zone of a sandy beach in northern France. Three experiments were executed to investigate (1) the effects of L. conchilega presence on sedimentary processes, as well as (2) the impacts of sedimentation on L. conchilega survival and patch growth, and (3) assess small-scale spatial heterogeneity in density and ecosystem engineering in L. conchilega aggregations. Weekly estimations of sedimentary properties in-situ showed that net deposition is significantly higher inside L. conchilega aggregations than in bare sand; whereas sediment mixing depth is noticeably reduced in comparison and regardless of patch tidal height. Variations in tube density above 3200 ind m-2 did not significantly impact sedimentary properties suggesting that the relationship between flow attenuation and tube density is nonlinear. In-situ monitoring of L. conchilega aggregations revealed different temporal trends for tube density and EPS content at the sediment surface between the center and edges of patches. This hints at the presence of environmental gradients within aggregations that may cause small-scale spatial heterogeneity. Finally, laboratory experiments showed significantly higher mortality rates and tube building activity in the presence of sediment deposition between 5 cm and 12 cm in column height. Results are in agreement with previous research suggesting that a positive feedback between sedimentation and tube-building activity drives the vertical expansion of patches. However, vertical expansion may be limited by deposition-induced mortality, thereby controlling population abundance.

Ecosystem engineers contribute to ecosystem functioning by regulating key environmental attributes, such as habitat availability and sediment biogeochemistry. While autogenic engineers can increase habitat complexity passively and provide physical protection to other species, allogenic engineers can regulate sediment oxygenation and biogeochemistry through bioturbation and/or bioirrigation. Their effects rely on the physical attributes of the engineer and/or its biogenic constructs, such as abundance and/or size. The present study focused on tube aggregations of a sessile, tube-building polychaete that engineers marine sediments, Lanice conchilega. Its tube aggregations modulate water flow by dissipating energy, influencing sedimentary processes and increasing particle retention. These effects can be influenced by temporal fluctuations in population demographic processes. Presently, we investigated the relationship between population processes and ecosystem engineering through an in-situ survey (1.5 years) of L. conchilega aggregations at the sandy beach of Boulogne-sur-Mer (France). We (1) evaluated temporal patterns in population structure, and (2) investigated how these are related to the ecosystem engineering of L. conchilega on marine sediments. During our survey, we assessed tube density, demographic structure, and sediment properties (surficial chl-a, EPS, TOM, median and mode grain size, sorting, and mud and water content) on a monthly basis for 12 intertidal aggregations. We found that the population was mainly composed by short-lived (6-10 months), small-medium individuals. Mass mortality severely reduced population density during winter. However the population persisted, likely due to recruits from other populations, which are associated to short- and long-term population dynamics. Two periods of recruitment were identified: spring/summer and autumn. Population density was highest during the spring recruitment and significantly affected several environmental properties (i.e. EPS, TOM, mode grain size, mud and water content), suggesting that demographic processes may be responsible for periods of pronounced ecosystem engineering with densities of approx. 30 000 ind m-2.

This paper focuses on the importance of biophysical interactions on short-term and long-term sediment dynamics. Therefore, various biological (macrobenthos, photopigments, colloidal EPS) and physical parameters (grain size, water content, sediment stability, bed level) were determined (bi)monthly in nine sampling plots on the IJzermonding tidal flat (Belgium, 51108 0

Distribution and structure of intertidal macrobenthic communities in the Zwin nature reserve, a lagoonal inlet consisting of marsh and tidal flat habitats, was investigated using univariate and multivariate analyses. Macrobenthos community structure was related to environmental characteristics and discussed in the framework of the implemented extension of the nature reserve.Based on explorative multivariate techniques, five different sample groups (SGs) were distinguished, which were, in general, located in different habitats of the Zwin nature reserve. The ecologically most important SGs consisted of the highest macrobenthic density, diversity and highest densities of Nereis diversicolor and Scrobicularia plana; these important prey species for wading birds and fish occurred in the deep tidal inlet channels. This habitat was characterized by fine to medium sand sediment and strong tidal currents, guaranteeing water renewal at each high tide. Other SGs were found in less and erratically submersed and thus stressed areas (i.e. tidal pond, salt pans and shallow flats). These assemblages were characterized by typical r-strategists (i.e. Capitella capitata and Polydora cornuta) and typical supralittoral, mobile species (i.e. Orchestia gammarellus and Collembola spp.). Being ecologically most important, the extension of wide, tidal creeks should be a prime target within the future development and management of the nature reserve.

The role of multi-species benthic diatom films (BDF) in the settlement of late pediveliger larvae of the bivalve Macoma balthica was investigated in still-water bioassays and multiple choice flume experiments. Axenic diatom cultures that were isolated from a tidal mudflat inhabited by M. balthica were selected to develop BDF sediment treatments characterized by a different community structure, biomass, and amount of extracellular polymeric substances (EPS). Control sediments had no added diatoms. Although all larvae settled and initiated burrowing within the first minute after their addition in still water, regardless of treatment, only 48-52% had completely penetrated the high diatom biomass treatments after 5 min, while on average 80 and 69% of the larvae had settled and burrowed into the control sediments and BDF with a low diatom biomass (3.5 µg Chl a g1 dry sediment), respectively. The percentage of larvae settling and burrowing into the sediment was negatively correlated with the concentration of Chl a and EPS of the BDF. This suggests higher physical resistance to bivalve penetration by the BDF with higher diatom biomass and more associated sugar and protein compounds. The larval settlement rate in annular flume experiments at flow velocities of 5 and 15cm s1 was distinctly lower compared to the still-water assays. Only 4.6-5.8% of the larvae were recovered from BDF and control sediments after 3h. Nonetheless, a clear settlement preference was observed for BDF in the flume experiments; i.e., larvae settled significantly more in BDF compared to control sediments irrespective of flow speed. Comparison with the settlement of polystyrene mimics and freeze-killed larvae led to the conclusion that active selection, active secondary dispersal and, at low flow velocities (5cm s1), passive adhesion to the sediment are important mechanisms determining the settlement of M. balthica larvae in estuarine biofilms.

The estuarine nitrogen cycle can be substantially altered due to anthropogenic activities resulting in increased amounts of inorganic nitrogen (mainly nitrate). In the past, denitrification was considered to be the main ecosystem process removing reactive nitrogen from the estuarine ecosystem. However, recent reports on the contribution of dissimilatory nitrate reduction to ammonium (DNRA) to nitrogen removal in these systems indicated a similar or higher importance, although the ratio between both processes remains ambiguous. Compared to denitrification, DNRA has been underexplored for the last decades and the key organisms carrying out the process in marine environments are largely unknown. Hence, as a first step to better understand the interplay between denitrification, DNRA and reduction of nitrate to nitrite in estuarine sediments, nitrogen reduction potentials were determined in sediments of the Paulina polder mudflat (Westerschelde estuary). We observed high variability in dominant nitrogen removing processes over a short distance (1.6 m), with nitrous oxide, ammonium and nitrite production rates differing significantly between all sampling sites. Denitrification occurred at all sites, DNRA was either the dominant process (two out of five sites) or absent, while nitrate reduction to nitrite was observed in most sites but never dominant. In addition, novel nitrate-to-ammonium reducers assigned to Thalassospira, Celeribacter, and Halomonas, for which DNRA was thus far unreported, were isolated, with DNRA phenotype reconfirmed through nrfA gene amplification. This study demonstrates high small scale heterogeneity among dissimilatory nitrate reduction processes in estuarine sediments and provides novel marine DNRA organisms that represent valuable alternatives to the current model organisms.

Surface sediments are important systems for the removal of anthropogenically derived inorganic nitrogen in estuaries. They are often characterized by the presence of a microphytobenthos (MPB) biofilm, which can impact bacterial communities in underlying sediments for example by secretion of extracellular polymeric substances (EPS) and competition for nutrients (including nitrogen). Pyrosequencing and qPCR was performed on two intertidal surface sediments of the Westerschelde estuary characterized by a two-fold difference in MPB biomass but no difference in MPB composition. Doubling of MPB biomass was accompanied by a disproportionately (ten-fold) increase in total bacterial abundances while, unexpectedly, no difference in general community structure was observed, despite significantly lower bacterial richness and distinct community membership, mostly for non-abundant taxa. Denitrifier abundances corresponded likewise while community structure, both for nirS and nirK denitrifiers, remained unchanged, suggesting that competition with diatoms for nitrate is negligible at concentrations in the investigated sediments (appr. 1 mg/l NO3-). This study indicates that MPB biomass increase has a general, significantly positive effect on total bacterial and denitrifier abundances, with stimulation or inhibition of specific bacterial groups that however do not result in a re-structured community.

Research so far has provided little evidence that benthic biogeochemical cycling is affected by ocean acidification under realistic climate change scenarios. We measured nutrient exchange and sediment community oxygen consumption (SCOC) rates to estimate nitrification in natural coastal permeable and fine sandy sediments under pre-phytoplankton bloom and bloom conditions. Ocean acidification, as mimicked in the laboratory by a realistic pH decrease of 0.3, significantly reduced SCOC on average by 60% and benthic nitrification rates on average by 94% in both sediment types in February (pre-bloom period), but not in April (bloom period). No changes in macrofauna functional community (density, structural and functional diversity) were observed between ambient and acidified conditions, suggesting that changes in benthic biogeochemical cycling were predominantly mediated by changes in the activity of the microbial community during the short-term incubations (14 days), rather than by changes in engineering effects of bioturbating and bio-irrigating macrofauna. As benthic nitrification makes up the gross of ocean nitrification, a slowdown of this nitrogen cycling pathway in both permeable and fine sediments in winter, could therefore have global impacts on coupled nitrification-denitrification and hence eventually on pelagic nutrient availability.

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Thresholds profoundly affect our understanding and management of ecosystem dynamics, but we have yet to develop practical techniques to assess the risk that thresholds will be crossed. Combining ecological knowledge of critical system interdependencies with a large-scale experiment, we tested for breaks in the ecosystem interaction network to identify threshold potential in real-world ecosystem dynamics. Our experiment with the bivalves Macomona liliana and Austrovenus stutchburyi on marine sandflats in New Zealand demonstrated that reductions in incident sunlight changed the interaction network between sediment biogeochemical fluxes, productivity, and macrofauna. By demonstrating loss of positive feedbacks and changes in the architecture of the network, we provide mechanistic evidence that stressors lead to break points in dynamics, which theory predicts predispose a system to a critical transition.

Habitat-modifying organisms that impact other organisms and local functioning are important in determining ecosystem resilience. However, it is often unclear how the outcome of interactions performed by key species varies depending on the spatial and temporal disturbance context which makes the prediction of disturbance-driven regime shifts difficult. We investigated the strength and generality of effects of the filter feeding cockle Cerastoderma edule on its ambient intertidal benthic physical and biological environment. By comparing the magnitude of the effect of experimental cockle removal between a non-cohesive and a sheltered cohesive sediment in two different periods of the year, we show that the outcome of cockle interference effects relates to differences in physical disturbance, and to temporal changes in suspended sediment load and ontogenetic changes in organism traits. Interference effects were only present in the cohesive sediments, though the effects varied seasonally. Cockle presence decreased only the density of surface-dwelling species suggesting that interference effects were particularly mediated by bioturbation of the surface sediments. Furthermore, density reductions in the presence of cockles were most pronounced during the season when larvae and juveniles were present, suggesting that these life history stages are most vulnerable to interference competition. We further illustrate that cockles may enhance benthic microalgal biomass, most likely through the reduction of surface-dwelling grazing species, especially in periods with high sediment load and supposedly also high bioturbation rates. Our results emphasize that the physical disturbance of the sediment may obliterate biotic interactions, and that temporal changes in environmental stressors, such as suspended sediments, may affect the outcome of key species interference effects at the local scale. Consequently, natural processes and anthropogenic activities that change bed shear stress and sediment dynamics in coastal soft-sediment systems will affect cockle-mediated influences on ecosystem properties and therefore the resilience of these systems to environmental change.

Bioturbation, the biogenic modification of sediments through particle reworking and burrow ventilation, is a key mediator of many important geochemical processes in marine systems. In situ quantification of bioturbation can be achieved in a myriad of ways, requiring expert knowledge, technology, and resources not always available, and not feasible in some settings. Where dedicated research programmes do not exist, a practical alternative is the adoption of a trait-based approach to estimate community bioturbation potential (BPc). This index can be calculated from inventories of species, abundance and biomass data (routinely available for many systems), and a functional classification of organism traits associated with sediment mixing (less available). Presently, however, there is no agreed standard categorization for the reworking mode and mobility of benthic species. Based on information from the literature and expert opinion, we provide a functional classification for 1033 benthic invertebrate species from the northwest European continental shelf, as a tool to enable the standardized calculation of BPc in the region. Future uses of this classification table will increase the comparability and utility of large-scale assessments of ecosystem processes and functioning influenced by bioturbation (e.g., to support legislation). The key strengths, assumptions, and limitations of BPc as a metric are critically reviewed, offering guidelines for its calculation and application.

This study investigated the effects of experimentally manipulated seawater carbonate chemistry on several early life history processes of the Baltic tellin (Macoma balthica), a widely distributed bivalve that plays a critical role in the functioning of many coastal habitats. We demonstrate that ocean acidification significantly depresses fertilization, embryogenesis, larval development and survival during the pelagic phase. Fertilization and the formation of a D-shaped shell during embryogenesis were severely diminished: successful fertilization was reduced by 11% at a 0.6 pH unit decrease from present (pH 8.1) conditions, while hatching success was depressed by 34 and 87%, respectively at a 0.3 and 0.6 pH unit decrease. Under acidified conditions, larvae were still able to develop a shell during the post-embryonic phase, but higher larval mortality rates indicate that fewer larvae may metamorphose and settle in an acidified ocean. The cumulative impact of decreasing seawater pH on fertilization, embryogenesis and survival to the benthic stage is estimated to reduce the number of competent settlers by 38% for a 0.3 pH unit decrease, and by 89% for a 0.6 pH unit decrease from present conditions. Additionally, slower growth rates and a delayed metamorphosis at a smaller size were indicative for larvae developed under acidified conditions. This may further decline the recruit population size due to a longer subjection to perturbations, such as predation, during the pelagic phase. In general, early life history processes were most severely compromised at ~pH 7.5, which corresponds to seawater undersaturated with respect to aragonite. Since recent models predict a comparable decrease in pH in coastal waters in the near future, this study indicates that future populations of Macoma balthica are likely to decline as a consequence of ongoing ocean acidification.

Hypoxia represents one of the major causes of biodiversity and ecosystem functioning loss for coastal waters. Since eutrophication-induced hypoxic events are becoming increasingly frequent and intense, understanding the response of ecosystems to hypoxia is of primary importance to understand and predict the stability of ecosystem functioning. Such ecological stability may greatly depend on the recovery patterns of communities and the return time of the system properties associated to these patterns. Here, we have examined how the reassembly of a benthic community contributed to the recovery of ecosystem functioning following experimentally-induced hypoxia in a tidal flat. We demonstrate that organism-sediment interactions that depend on organism size and relate to mobility traits and sediment reworking capacities are generally more important than recovering species richness to set the return time of the measured sediment processes and properties. Specifically, increasing macrofauna bioturbation potential during community reassembly significantly contributed to the recovery of sediment processes and properties such as denitrification, bedload sediment transport, primary production and deep pore water ammonium concentration. Such bioturbation potential was due to the replacement of the small-sized organisms that recolonised at early stages by large-sized bioturbating organisms, which had a disproportionately stronger influence on sediment. This study suggests that the complete recovery of organism-sediment interactions is a necessary condition for ecosystem functioning recovery, and that such process requires long periods after disturbance due to the slow growth of juveniles into adult stages involved in these interactions. Consequently, repeated episodes of disturbance at intervals smaller than the time needed for the system to fully recover organism-sediment interactions may greatly impair the resilience of ecosystem functioning.

The distribution and general population characteristics of five mysid shrimps were investigated in the period 2003–2005 in the Westerschelde estuary, a tidal temperate estuary situated along the Dutch–Belgian border. Multivariate analyses revealed that salinity predominantly governs the spatial distribution of Neomysis integer, Gastrosaccus spinifer, Schistomysis kervillei and Schistomysis spiritus while temperature, and to a lesser extent turbidity, control the distribution of Mesopodopsis slabberi. N. integer is a resident species in the mesohaline zone of the estuary, i.e. all life stages of the species are present in the estuary throughout the year. For the first time since decades N. integer inhabits the oligohaline zone of the estuary supposedly as a consequence of improved oxygen conditions in the upstream reaches. M. slabberi, the most abundant mysid in the Westerschelde, dominates the hyperbenthos of the mesohaline zone of the estuary. The polyhaline zone of the estuary is, most abundantly inhabited by M. slabberi, G. spinifer, S. kervillei and to a lesser extent by S. spiritus. The abundance of the latter four species is low in winter, probably due to a migration towards coastal waters to avoid colder temperatures in the estuary combined with an increased mortality after breeding. The sex ratio of all the mysid populations corresponds to the expected 1:1 female:male ratio and no salinity governed segregation is found between the different life stages of each mysid population. A seasonal variation exists in brood size in the N. integer population regardless of the body size, with a larger number of broods during winter and spring compared to the summer. In the other mysid populations the brood sizes vary only with the length of the ovigerous females.Our recent observations underline some general characteristics of mysid populations in the Westerschelde and provide novel insights in their life stage and sex specific population segregation, their brood sizes and the expanding upstream distribution of N. integer.